Gecarcinus ruricola
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''Gecarcinus ruricola'' is a
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of
terrestrial crab A number of lineages of crabs have evolved to live predominantly on land. Examples of terrestrial crabs are found in the families Gecarcinidae and Gecarcinucidae, as well as in selected genera from other families, such as '' Sesarma'', althoug ...
. It is the most terrestrial of the Caribbean land crabs, and is found from western
Cuba Cuba ( , ), officially the Republic of Cuba ( es, República de Cuba, links=no ), is an island country comprising the island of Cuba, as well as Isla de la Juventud and several minor archipelagos. Cuba is located where the northern Caribbea ...
across the
Antilles The Antilles (; gcf, label=Antillean Creole, Antiy; es, Antillas; french: Antilles; nl, Antillen; ht, Antiy; pap, Antias; Jamaican Patois: ''Antiliiz'') is an archipelago bordered by the Caribbean Sea to the south and west, the Gulf of Mex ...
as far east as
Barbados Barbados is an island country in the Lesser Antilles of the West Indies, in the Caribbean region of the Americas, and the most easterly of the Caribbean Islands. It occupies an area of and has a population of about 287,000 (2019 estimate) ...
. Common names for ''G. ruricola'' include the purple land crab, black land crab, red land crab, and zombie crab.


Description

Four colour morphs exist within the species - black, red, yellow, and green. The carapace of ''G. ruricola'' grows in width at a rate of about per year, with the crabs reaching maturity after 5 years, and living for up to 10 years in total. ''G. ruricola'' crabs have a number of adaptations to terrestrial life, mostly regarding
water conservation Water conservation includes all the policies, strategies and activities to sustainably manage the natural resource of fresh water, to protect the hydrosphere, and to meet the current and future human demand (thus avoiding water scarcity). Popula ...
. They are nocturnal, to prevent the hot sun from drying them out. They also have a "nephritic pad", onto which
urine Urine is a liquid by-product of metabolism in humans and in many other animals. Urine flows from the kidneys through the ureters to the urinary bladder. Urination results in urine being excreted from the body through the urethra. Cellular ...
is released, to be cleaned by microbes before the water is then reabsorbed.


Distribution

''G. ruricola'' is found across much of the Caribbean, from Cuba and the
Bahamas The Bahamas (), officially the Commonwealth of The Bahamas, is an island country within the Lucayan Archipelago of the West Indies in the North Atlantic. It takes up 97% of the Lucayan Archipelago's land area and is home to 88% of the ar ...
in the west through the Antilles to Barbados in the east. It has been reported from
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
and
Nicaragua Nicaragua (; ), officially the Republic of Nicaragua (), is the largest country in Central America, bordered by Honduras to the north, the Caribbean to the east, Costa Rica to the south, and the Pacific Ocean to the west. Managua is the countr ...
, but few confirmed examples exist from the mainland;
Loggerhead Key Loggerhead Key is an uninhabited tropical island within the Dry Tortugas group of islands in the Gulf of Mexico. At approximately 49 acres (19.8 hectares) in size, it is the largest island of the Dry Tortugas. Despite being uninhabited, the is ...
in the
Dry Tortugas Dry Tortugas National Park is a national park located about west of Key West in the Gulf of Mexico. The park preserves Fort Jefferson and the seven Dry Tortugas islands, the westernmost and most isolated of the Florida Keys. The archipelago's c ...
marks the northernmost limit of its island distribution, which extends across the Bahamas and Cuba, through the Greater and Lesser Antilles, to Barbados. Outlying populations exist on Curaçao, in the Swan Islands off Honduras, Half Moon Caye of Belize, and the
Archipelago of San Andrés, Providencia and Santa Catalina The Archipelago of San Andrés, Providencia and Santa Catalina ( es, Archipiélago de San Andrés, Providencia y Santa Catalina, ), or San Andrés and Providencia, is one of the departments of Colombia, and the only one in North America. It consis ...
off the Colombian coast. It can be found at great distances from the sea, and at high
altitude Altitude or height (also sometimes known as depth) is a distance measurement, usually in the vertical or "up" direction, between a reference datum and a point or object. The exact definition and reference datum varies according to the context ...
s; it has been observed above on the island of Dominica, and at over on
Jamaica Jamaica (; ) is an island country situated in the Caribbean Sea. Spanning in area, it is the third-largest island of the Greater Antilles and the Caribbean (after Cuba and Hispaniola). Jamaica lies about south of Cuba, and west of His ...
.


Life cycle

The
eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
of ''G. ruricola'' hatch in the sea, where the
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
e live as
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucia ...
. The return of the larvae to land seems to be infrequent, but when they do return, they return as
megalopa Crustaceans may pass through a number of larval and immature stages between hatching from their eggs and reaching their adult form. Each of the stages is separated by a moult, in which the hard exoskeleton is shed to allow the animal to grow. The ...
larvae, in sufficient numbers to turn roads red. They move at speeds of , or faster if startled. For the next three years, the young crabs live in burrows inhabited by other crabs, and eat food brought back to the burrow by the older crab. After mating, mass migrations occur, with the females returning to the sea to release their fertilised eggs. A typical female carries around 85,000 eggs.


Ecology

''G. ruricola'' is an
omnivorous An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nut ...
scavenger, feeding mostly on
nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
-poor plant matter. The meat of ''G. ruricola'' is rich in
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
, and has often been harvested by local people. Other
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s may include
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s, although information is scarce. When confronted, they rear up and hold their open claws outwards in a defensive posture.


''Drosophila''

''G. ruricola'' is the
host organism In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasitic, a mutualistic, or a commensalist ''guest'' (symbiont). The guest is typically provided with nourishment and shelter. Examples include a ...
for two species of
commensal Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit fro ...
flies Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced m ...
in the genus ''
Drosophila ''Drosophila'' () is a genus of flies, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or (less frequently) pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many speci ...
''. They were first observed on Montserrat by Henry Guernsey Hubbard in 1894, and presented at a
scientific meeting An academic conference or scientific conference (also congress, symposium, workshop, or meeting) is an event for researchers (not necessarily academics) to present and discuss their scholarly work. Together with academic or scientific journals a ...
later that year (where the crab was misidentified as ''
Cardisoma guanhumi ''Cardisoma guanhumi'', also known as the blue land crab, is a species of land crab found in tropical and subtopical estuaries and other maritime areas of land along the Atlantic coast of the Americas from Brazil and Colombia, through the Ca ...
''), but no further research was conducted until 1955, when specimens were again collected, this time from
Mona Island Mona ( es, Isla de Mona) is the third-largest island of the Puerto Rican archipelago, after the main island of Puerto Rico and Vieques. It is the largest of three islands in the Mona Passage, a strait between the Dominican Republic and Puerto R ...
, and named as ''Drosophila carcinophila'' by M. R. Wheeler. In 1967, a second species of fly, '' Drosophila endobranchia'', was discovered on ''G. ruricola'', although it also inhabits the closely related species '' G. lateralis''. ''D. endrobranchia'' is not closely related to ''D. carcinophila'', and this trait appears to have evolved convergently. (A third species of fly, '' Lissocephala powelli'', has evolved a similar habit on
Christmas Island Christmas Island, officially the Territory of Christmas Island, is an Australian external territory comprising the island of the same name. It is located in the Indian Ocean, around south of Java and Sumatra and around north-west of the ...
, where it lives on the land crabs '' Gecarcoidea lalandii'', '' Geograpsus crinipes'', and '' Cardisoma carnifex'', and the terrestrial
hermit crab Hermit crabs are anomuran decapod crustaceans of the superfamily Paguroidea that have adapted to occupy empty scavenged mollusc shells to protect their fragile exoskeletons. There are over 800 species of hermit crab, most of which possess an as ...
'' Birgus latro''.) ''D. endobranchia'' evolved from a group of species that breeds on fungi or bark, while ''D. carcinophila'' evolved from a group that breeds on cacti. The flies spend most of their lives on the crab, and are reluctant to leave. They do not need to flee predators, because the crabs they inhabit are fast animals and will flee:
The flies … hardly move at all, are extremely reluctant in leaving their host crabs and are hard pushed to take flight. Although the flies are sluggish, the crabs on which they reside are anything but. Chasing after crabs through a pitch-black jungle (growing on a razor-sharp labyrinthine limestone ground), while trying to aspirate flies from their carapaces is not trivial. Obtaining large amounts of flies in this way is simply a nightmare.
The
eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
are laid around the crab's
compound eye A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which disti ...
s; the first-
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ...
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
e migrate to the crab's nephritic pad, and live there, feeding on the microbes that cleanse the crab's urine. The second instar is spent in the crab's
gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
chamber. The third instar has the larvae return to the mouthparts before falling to the ground to
pupate A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
.


Linnaeus

Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his Nobility#Ennoblement, ennoblement in 1761 as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalise ...
described the species in 1758 (the starting point for
zoological nomenclature The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its publisher, the In ...
), noting the species' annual migrations from the forests to the coast (''Habitat in America, sylvas vastissimis agminibus quotannis deserens littora maris petiturus'': "lives in America; every year, an army marches out of the forests towards the sea").


Philately

''G. ruricola'' appeared on two
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
n
postage stamp A postage stamp is a small piece of paper issued by a post office, postal administration, or other authorized vendors to customers who pay postage (the cost involved in moving, insuring, or registering mail), who then affix the stamp to the f ...
s for the ''International Year of the Ocean'' in 1998, under the name "mountain crab". These were a
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands ...
n stamp worth TSh  and a
Uganda }), is a landlocked country in East Africa. The country is bordered to the east by Kenya, to the north by South Sudan, to the west by the Democratic Republic of the Congo, to the south-west by Rwanda, and to the south by Tanzania. The sou ...
n stamp worth USh .


References

{{Taxonbar, from=Q5233916 Grapsoidea Terrestrial crustaceans Edible crustaceans Crustaceans of the Atlantic Ocean Crustaceans described in 1758 Arthropods of the Dominican Republic Taxa named by Carl Linnaeus